''Boquila'' is a
monotypic
In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
genus
Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s in the family
Lardizabalaceae
Lardizabalaceae is a family of flowering plants.
The family has been universally recognized by taxonomists, including the APG II system (2003; unchanged from the APG system of 1998), which places it in the order Ranunculales, in the clade eudic ...
, native to temperate forests of central and southern
Chile
Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the east a ...
and
Argentina
Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. The sole species is ''Boquila trifoliolata'' (DC.) Decne., called ''pilpil, voqui, voquicillo, voquillo,'' and ''voqui blanco'' in Chile. It grows vines that wrap around host plants, mimicking the host's leaves in a phenomenon called mimetic polymorphism. It bears an edible fruit (Boquila berries).
This species has been shown to be capable of
mimicking the leaves of its supporting trees.
Ernesto Gianoli said "Boquila's leaves are extraordinarily diverse. The biggest ones can be 10 times bigger than the smallest, and they can vary from very light to very dark. In around three-quarters of cases, they're similar to the closest leaf from another tree, matching it in size, area, length of stalk, angle, and color. Boquila's leaves can even grow a spiny tip when, and only when, it climbs onto a shrub with spine-tipped leaves." When there are no nearby host leaves to influence them, the normal leaves of ''B. trifoliolata'' are relatively short and light green with rounded edges.
''Boquila'', unlike other plants capable of mimicry, does not require physical contact to match its host.
Mimicry
''Boquila trifoliolata'' is unique because of the ability of its leaves to mimic those of the hosts that are supporting them, a phenomenon called mimetic polymorphism. This plant's climbing behavior protects it from ground-dwelling herbivores, and leaf mimicry is a protection against leaf herbivores. ''B. trifoliolata'' differs from other plants that can mimic a host, like the Australian
mistletoe
Mistletoe is the common name for obligate hemiparasitic plants in the order Santalales. They are attached to their host tree or shrub by a structure called the haustorium, through which they extract water and nutrients from the host plant.
...
, because it is not limited to mimicking a single host and because it is not a parasite to the host tree. An individual ''B. trifoliolata'' vine can mimic multiple kinds of foliage that are closest in proximity to it.
Their mimicking behavior was discovered by researchers Ernesto Gianoli and Fernando Carrasco-Urra. They carried out observations and measurements in a rainforest located at
Puyehue National Park
Puyehue National Park () is located in the Andes mountain range, in Los Ríos and Los Lagos regions of Chile formerly referred to as the 10th region. The park boast 220,000 acres of natural thermal springs, volcanoes, and evergreen forests, afte ...
in southern
Chile
Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the east a ...
. They sampled 12 different species of host trees with 45 total individual ''B. trifoliolata'' vines that had climbed these trees. The two closest leaves in proximity between a pair of the 45 vine-trees were measured, 11 different traits in total: angle, thickness, petiole length, leaflet petiole length, leaflet angle, maximum width, maximum length, area, perimeter, area/perimeter, and color. Usage of a
generalized linear model
In statistics, a generalized linear model (GLM) is a flexible generalization of ordinary linear regression. The GLM generalizes linear regression by allowing the linear model to be related to the response variable via a ''link function'' and b ...
showed that 9 of the 11 traits demonstrated mimicry by the vine to its host tree. Gianoli et al. also sampled more individuals that were prostrated, that grew on leafless tree trunks, and more individuals that have climbed on the 8 most common host species. To analyze these samples, the researchers used
multivariate analysis of variance
In statistics, multivariate analysis of variance (MANOVA) is a procedure for comparing multivariate sample means. As a multivariate procedure, it is used when there are two or more dependent variables, and is often followed by significance tests i ...
(MANOVA). They found that the prostrate individuals were not different from the leafless-host vines, but that they were different for 7 of the 8 common-host vine leaves. They also concluded that the leafless-host vines were different for 6 of the 8 common-host vines. Moreover, this leaf mimicry served as a protection against leaf herbivores and led to lower leaf herbivory rates. Climbing vines had no difference in herbivory compared to supporting host tree leaves but had much lower herbivory compared to prostrated, unsupported ''B. trifoliolata'' individuals. The highest amount of herbivory was on ''B. trifoliolata'' vines that climbed onto leafless hosts.
Currently, there is no known mechanism for how ''B. trifoliolata'' is able to mimic host leaves so well, but Gianoli proposes two possible mechanisms. One hypothesis is that
volatile organic compounds
Volatile organic compounds (VOCs) are organic compounds that have a high vapour pressure at room temperature
Colloquially, "room temperature" is a range of air temperatures that most people prefer for indoor settings. It feels comfortable to a ...
emitted from host plant leaves induce a phenotypic change in nearby ''B. trifoliolata'' leaves. By receiving different host signals into its system, it is able to create specific signals and hormones in its tissues to regulate gene transcription of leaf morphology and developmental pathways for leaf differentiation. The other hypothesis is that there could be
horizontal gene transfer
Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between Unicellular organism, unicellular and/or multicellular organisms other than by the ("vertical") transmission of DNA from parent to offsprin ...
between the host and ''B. trifoliolata''. A separate study also conducted by Gianoli et al. suggests that bacterial agents, which could mediate a horizontal gene transfer, may play a role in leaf mimicry by ''B. trifoliolata''. A comparison of
endophytic
An endophyte is an endosymbiont, often a bacterium or fungus, that lives within a plant for at least part of its life cycle without causing apparent disease. Endophytes are ubiquitous and have been found in all species of plants studied to date; h ...
bacterial communities among leaves from a host tree, ''B. trifoliolata'' leaves mimicking the host tree leaves, and non-mimetic ''B. trifoliolata'' leaves from the same vine revealed that the bacterial communities were more similar among mimetic ''B. trifoliolata'' and host leaves than among non-mimetic ''B. trifoliolata'' and host leaves.
In a 2021 paper, it was suggested that the plant has some sort of vision using
ocelli
A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
. This hypothesis was presented on the basis of experiments in which the vine appeared to mimic plastic vines and artificial plants. However, the only control used was the lower leaves on the same plants, with an opaque shelf over them which could have influenced light exposure. The lack of control plants climbing anything apart from the same model of artificial host plant makes it plausible that the observed differences were due to age and light exposure.
See also
* ''
Lardizabala
''Lardizabala'' is a monotypic genus of flowering plants. These plants are evergreen lianas, native to temperate forests of central and southern Chile.
The sole species is ''Lardizabala biternata'' Ruiz & Pav, known as ''Coguil, Cogüilera, Coi ...
'', a related species also grown for its fruit
References
*
*
{{Taxonbar, from=Q3753460, from2=Q15601133
Lardizabalaceae
Flora of Chile
Monotypic Ranunculales genera
Taxa named by Joseph Decaisne